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POLLEN MORPHOLOGY of KALMIA L. (PHYLLODOCEAE, ERICACEAE) and ITS TAXONOMIC Title SIGNIFICANCE

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POLLEN MORPHOLOGY of KALMIA L. (PHYLLODOCEAE, ERICACEAE) and ITS TAXONOMIC Title SIGNIFICANCE POLLEN MORPHOLOGY OF KALMIA L. (PHYLLODOCEAE, ERICACEAE) AND ITS TAXONOMIC Title SIGNIFICANCE Author(s) Sarwar, A.K.M.Golam; Takahashi, Hideki Citation Bangladesh Journal of Plant Taxonomy, 19(2), 123-133 Issue Date 2012-12 Doc URL http://hdl.handle.net/2115/51240 Type article File Information BJPT19_123.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP Bangladesh J. Plant Taxon. 19(2): 123-133, 2012 (December) © 2012 Bangladesh Association of Plant Taxonomists POLLEN MORPHOLOGY OF KALMIA L. (PHYLLODOCEAE, ERICACEAE) AND ITS TAXONOMIC SIGNIFICANCE 1 A.K.M. GOLAM SARWAR AND HIDEKI TAKAHASHI Laboratory of Systematic Botany, Graduate School of Agriculture, Hokkaido University, Japan Keywords: Pollen morphology; Infrageneric classification; Exine sculpture; Kalmia. Abstract Pollen morphology of seven taxa of Kalmia was examined using light and scanning electron microscopy (LM and SEM, respectively), or SEM alone, in search of new characters that might contribute to infrageneric classification of the genus. The Kalmia species are stenopalynous and characterized by 3-colpor(oid)ate, medium, oblate pollen united in tetrahedral tetrads with rugulate exine sculpture. However, a continuous and serial variation in all the quantitative characters and exine sculpture was revealed within the genus. Kalmia buxifolia is characterized by having the smallest pollen tetrads, largest aperture, largest 2f/D ratio and septum thicker than apocolpial exine. On the other hand, K. latifolia produces the largest pollen tetrads with smallest aperture and smallest 2f/D ratio. An evolutionary trend from rugulate to psilate has also been observed in apocolpial exine sculpture of the genus Kalmia. On the basis of acetolysed pollen characteristics, a dichotomous key for Kalmia was prepared. Introduction Kalmia L. (Family Phyllodoceae, subfamily Ericaceae) comprises eleven quite distinct species of evergreen or deciduous shrubs (rarely tree) and is restricted to North America and Cuba, the only circumboreal species is K. procumbens (Kron et al., 2002; Stevens et al., 2004). The number of species (including Leiophyllum Pers. and Loiseleuria Desv.) varies from seven to eleven (Ebinger, 1974; Southall and Hardin, 1974; Judd, 1995; Stevens et al., 2004). Linnaeus described the genus Kalmia (Linnaeus, 1753, 1754) as well as its first two species, K. latifolia and K. angustifolia. Drude (1889) placed Kalmia in the tribe Phyllodoceae, and its position has not been changed by most of the subsequent authors (see Ebinger, 1974). There have been few nomenclatural complications in the genus, and only two other generic names have been proposed. Kuntze (1891) used Catesby's pre-Linnean name of Chamaedaphne and made numerous transfers to it, while Small (1914) divided Kalmia into two genera. He proposed the genus Kalmiella, segregating Kalmia hirsuta and K. ericoides in the latter genus on the basis of their inflorescence structure (i.e., solitary flowers) and deciduous calyx (Small, 1914). All subsequent authors (Copeland, 1943; Wood, 1961), except Alain (1957), have treated Kalmiella as a synonym of Kalmia. Leiophyllum Pers. and Loiseleuria Desv. are two monotypic genera of the tribe Phyllodoceae, Ericaceae (Drude,1889; Stevens, 1971). Leiophyllum buxifolia (Berg.) Elliott is a shrub endemic to the southeastern United States (New Jersey). Due to its disjunct distribution and morphological variability, Leiophyllum has long been a subject of debate and speculation about its taxonomy and evolutionary history (Strand and Wyatt, 1991). Loiseleuria procumbens (L.) Desv. has Holarctic circumpolar-boreal range with a large gap in North-West Asia. Loiseleuria and Leiophyllum despite differences in the leaf vernation and fusion of corolla, show many similarities in anatomy 1 Present Address: Department of Crop Botany, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh. E-mail: [email protected] 124 SARWAR AND TAKAHASHI and anther structure, and described as a closely related pair of genera (Stevens, 1971). Recent molecular phylogenetic studies included these two monotypic genera within Kalmia (Kron and King, 1996; Kron et al., 2002). Here, we adopt a broad circumscription of Kalmia which includes Leiophyllum and Loiseleuria [see Kron et al. (2002) for the formal transfers]. Pollen of Kalmia is shed in tetrahedral tetrads which vary in size from 45 µm in K. latifolia to 23 µm in K. angustifolia var. carolina; the tetrads are associated with viscin strands in K. latifolia, K. hirsuta, K. ericoides and K. angustifolia, while such threads have not been seen in the remaining species (Ebinger, 1974; Southall and Hardin, 1974; Judd, 1995). Exine sculpturing is generally coarsely reticulate or rugulate in K. microphylla (Hook.) Heller, while finely so in K. latifolia and K. ericoides. The pollen morphology of some taxa has been described in previous literature, but little is known about the use of palynological features in taxonomy/systematics. Hitherto, no formal infrageneric classification of Kalmia has been proposed, but some distinctly different groups, based on different features, have been recognized within this genus (Stevens, 1971; Ebinger, 1974; Southall and Hardin, 1974; Judd, 1995; Kron and King, 1996). Previously, the pollen morphological features were found to be important in the infrageneric as well as subfamilial classification of some Ericaceous taxa (Sarwar and Takahashi, 2006a, b, 2009; Sarwar et al., 2006; Sarwar, 2011). Therefore, we present here a general pollen survey on Kalmia based on LM and SEM to search new character that could add information pertinent to infrageneric classification of this genus. Materials and Methods Pollen morphology of seven taxa (and 11 specimens) out of 11 species of Kalmia was examined by means of LM and SEM, or SEM alone (Table 1). The remaining four species were not examined due to unavailability of herbarium specimens (and/or pollen samples) in the herbaria consulted here as well as others (Sarwar, 2011; for the complete list of herbaria consulted). Polliniferous materials used in this investigation were taken from the dried specimens from the herbaria S, SAPS and SAPT. Abbreviation of the herbarium names except for SAPT (the Botanic Garden, Hokkaido University, Sapporo) are according to the Index Herbariorum (Holmgren et al., 1990). Pollen grains were acetolysed following the technique of Erdtman (1960) modified by Takahashi (1987). Palynological features of three Kalmia species were studied both acetolysed and non-acetolysed condition to find out the effect of acetolysis on its pollen. For LM, the dehydrated (in an ethanol series) pollen (both acetolysed and non-acetolysed) was mounted in silicone oil (viscosity 3000 cs), and examined and measured with a Nikon Eclipse E200 microscope. The dimensions “D”, “P”, “d (E)” and “2f”, corresponding to the tetrad diameter, polar length, equatorial length and colpus length of pollen grain were measured, and the D/d, P/E and 2f/D ratio was calculated (Oldfield, 1959). The arithmetic mean, standard deviation and the maximum and minimum values were calculated using the XLSTAT 2009.3 program. The measurements given in Table 2 are based on at least 10 grains from each specimen. Principal component analysis (PCA) and agglomerative hierarchical clustering (AHC) were also conducted using the same program to visualise the relationships among the studied species based on pollen data. A dendrogram was built by AHC. Pollen slides of all collections are deposited at the Hokkaido University Museum, Sapporo, Japan. Pollen size and shape classes were made following Erdtman (1986) and descriptive terminology follows Punt et al. (2007) and Sarwar et al. (2006). For SEM, the pollen samples, both acetolysed and non-acetolysed, were dehydrated in an ethanol series, and mounted and air dried on aluminum stubs from 70% ethanol, and sputter coated POLLEN MORPHOLOGY OF KALMIA L. 125 with Platinum-Palladium by a Hitachi E102 ion sputter. Subsequently, these were examined and photographed with a Jeol JSM-5310 LV scanning electron microscope operated at 15 KV. The SEM micrographs of apocolpial exine sculpture of similar position were used for the purpose of description and comparison. Table 1. List of Kalmia taxa used in this study along with their voucher specimens. No. Taxa Voucher specimens 1. Kalmia angustifolia L. USA: Connecticut, Litchfield Co., Norfolk, Great Mt. Forest, Tobey bog, 23.06.1983. S. DeSimon 415 (SAPT) North America: Grand Harbor, 03.08.1888, K. Miyabe s.n. (SAPS) 2. K. buxifolia (Berg.) Gift, Kron & Stevens USA: North Carolina, Brunswi. Co., 12 miles S of Wilmington, 06.04.1939. R.K. Godfrey & R.N. White 7110 (S) 3. K. ericoides Wright ex Grisebatch var. Cuba: Prov. Pinar del Rio, La Grifa, 19.11.1923. E.L. aggregata (Small) Ebinger Ekman 18165 (S) 4. K. latifolia L. USA: Connecticut, Litchfield Co., Mt. Riga State Park, near Riga lake, moist woods, 16.06.1981. G. Feldman 63 (SAPT) Georgia, Lumpkin Co., Chattahoochee National Park, 11.05.1982, F.H. Utech et al. 82-056 (SAPS) 5. K. microphylla (Hook.) Heller var. USA: Washington, Mt. Rainier, (Flora of Cascade microphylla Mountains), 13.08.1894. O.D. Allen s.n. (SAPS) Canada: British Columbia, Garibaldi Park, 23.07.1975, V.J. Krajina s.n. (SAPT) 6. K. polifolia Wangenh. USA: Washington, swamp, upper valley of the Nesqually, (Flora of Cascade Mountains), 27.05.1893. O.D. Allen s.n. (SAPS) 7. K. procumbens (L.) Gift, Kron & Stevens Japan: Hokkaido, Prov. Kitami, Monbestu-gun, Shirataki-mura, Mt. Taira-yama, 30.06.1980. H. Takahashi et al. 2644 (SAPS) Sweden: Torne Lappmark, Abisco, Mt Njulla, 20.07.1989. H. Takahashi 9907 (SAPS) Results General pollen morphology In LM, the medium pollen grains are united in tetrahedral tetrad (Fig. 1A-C); viscin threads present in some species; D 24.4-36.6 µm, P 12.8 -18.8 µm, E 17.3-26.3 µm, D/d 1.31-1.52, P/E 0.70-0.80, oblate or suboblate; 3-colpor(oid)ate, 2f 11.6-19.6 µm, W 0.4-1.2 µm, 2f/D 0.32-0.67, costae present and distinct, colpus margin distinct; endocracks present; endoaperture distinct and lalongate; apocolpial exine 1.6-2.1 µm thick, septum 0.7-1.9 µm thick (Table 2); tectate, apocolpial exine sculpture from fine verrucate to rugulate.
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